The circle of Willis revisited: Forebrain dehydration sensing facilitated by the anterior communicating artery: How hemodynamic properties facilitate more efficient dehydration sensing in amniotes.

We hypothesize that threat of dehydration provided selection pressure for the evolutionary emergence and persistence of the anterior communicating artery (ACoA - the inter-arterial connection that completes the Circle of Willis) in early amniotes. The ACoA is a hemodynamically insignificant artery, but, as we argue in this paper, its privileged position outside the blood-brain barrier gives it a crucial sensing function for the osmolarity of the blood against the background of the rest of the brain, which efficiently protects itself from dehydration. Till now, the questions of why the ACoA evolved, and what its physiological function is, have remained unsatisfactorily answered. The traditional view-that the ACoA serves as a collateral source of vascularization in case of arterial stenosis-is anthropocentric, and not in accordance with principles of natural selection that apply more generally. Diseases underlying arterial stenosis are associated with aging and the human lifestyle, so this cannot explain why the ACoA formed hundreds of millions of years ago and persisted in amniotes to this day. The peculiar hemodynamic properties of the ACoA could be selected traits that allowed for more efficient forebrain detection of dehydration and complex behavioral responses to water loss, a major advantage in the survival of early amniotes. This hypothesis also explains insufficient hydration often seen in elderly humans.